The Ultramafic Flora of the Santa Elena Peninsula, Costa Rica: a Biogeochemical Reconnaissance ⁎ R.D
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Journal of Geochemical Exploration 93 (2007) 153–159 www.elsevier.com/locate/jgeoexp The ultramafic flora of the Santa Elena peninsula, Costa Rica: A biogeochemical reconnaissance ⁎ R.D. Reeves a, , A.J.M. Baker a, R. Romero b a School of Botany, University of Melbourne, Australia b Department of Chemistry, University of Costa Rica, San Jose, Costa Rica Received 16 October 2006; accepted 9 April 2007 Available online 19 April 2007 Abstract The Santa Elena peninsula in the northwest of Costa Rica protrudes about 30 km westwards into the Pacific Ocean, and measures about 8–16 km in a north–south direction. Several geological studies have been carried out since 1953, showing that much of the peninsula is made up of peridotite, cut by mafic dykes. Only one previous brief examination appears to have been made of the vegetation in relation to the composition of the soils. We present here the results of a survey of some soils and plants of the eastern part of the peridotite massif, in which 73 plant specimens representing 51 identified species were collected and analyzed. The soils sampled all showed extreme ultramafic characteristics: Fe 10–16%, Mg 4–16%, Ca 0.5–1.4%; Ni 3000– 7500 mg/kg, Cr 1400–3650 mg/kg, Co 150–325 mg/kg. The plants collected include several from genera such as Arrabidaea, Chamaesyce, Helicteres, Hyptis, Lippia, Oxalis, Polygala, Turnera and Waltheria that are also represented on ultramafics elsewhere in the Americas (e.g. Cuba, Puerto Rico, Brazil). Few of the species appear to be endemic to Costa Rica or to the ultramafics of Santa Elena. None of the specimens collected exhibited hyperaccumulation of nickel, the highest Ni concentration being 275 mg/kg in Buchnera pusilla. © 2007 Elsevier B.V. All rights reserved. Keywords: Costa Rica; Ultramafic flora; Serpentine; Nickel; Biogeochemistry; Santa Elena 1. Introduction silicate mineral serpentine in many ultramafics) have been extensively studied in many parts of the world Soils derived from ultramafic rocks, which are rich in during the last hundred years, leading to a wealth of iron, magnesium and associated trace elements such as knowledge about the limited range of plant species that nickel, cobalt and chromium, have for many years been are able to tolerate the unusual chemical environment of particular interest to scientists working in fields these soils provide. Not only are these soils character- where geochemistry and botany intersect. Ultramafic ized by elevated levels of elements that may be toxic, floras (sometimes referred to as ‘serpentine’ floras, from but the levels of important plant nutrients such as the frequently important presence of the magnesium calcium, potassium and phosphorus are generally unusually low, providing an additional stress factor for plant survival and growth. ⁎ Corresponding author. During the last two decades, several comprehensive E-mail address: [email protected] (R.D. Reeves). publications with worldwide scope (Brooks, 1987; 0375-6742/$ - see front matter © 2007 Elsevier B.V. All rights reserved. doi:10.1016/j.gexplo.2007.04.002 154 R.D. Reeves et al. / Journal of Geochemical Exploration 93 (2007) 153–159 Roberts and Proctor, 1992; Baker et al., 1992) have dealt peridotites are dominated by diopside-bearing harzbur- with many aspects of serpentine floras: plant inventories gites–lherzolites typically containing 0.5–1.0% Cr2O3 and taxonomy, plant geography, evolution of endemic and 0.2–0.5% NiO, and are richer in Al and Ca than is species, physiology of metal tolerance, and the occa- usual in harzburgites (Desmet, 1985). The geology of sional occurrence of extreme nickel accumulation to this area has been discussed in some detail by Harrison levels exceeding 1000 mg/kg in the plant dry matter (1953), Jager (1977), de Boer (1979), Azéma and (‘hyperaccumulation’). Other publications describe Tournon (1980), Desmet (1985) and Tournon (1994), work carried out on ultramafic floras of a single country amongst others. or region, e.g. New Caledonia (Jaffré, 1980), California Botanical work on the peninsula up to the present (Kruckeberg, 1984), Cuba (Borhidi, 1991). However, time appears to be largely confined to the ongoing there are still large gaps in our knowledge of the species preparation of species inventories for the Parque composition and chemical behavior of serpentine plants Nacional Santa Rosa, and to an unpublished study in many parts of the world, such as in Central America, from the Río Murciélago area (sedimentary–ultramafic Brazil, the Philippines and Indonesia, and there is a need boundary several km north of the area studied here) for more plant collection, taxonomic work and chemical (Balding, 1993). The latter work compared vegetation of analysis of plants and their associated soils from these several 1-m2 quadrats on mudstone with several on areas. serpentine less than 2 km away. Its value is severely One of the two major ultramafic areas of Costa Rica limited by the fact that only 10 of the species found were is on the Santa Elena peninsula on the Pacific coast near identified at the species level. the border with Nicaragua. The peninsula measures Part of the interest in the present study lay in trying to about 30 km E–W and 8–16 km N–S, and falls within determine whether the ultramafics of Santa Elena the Parque Nacional Santa Rosa (approx. 10°40′– supported a significant number of endemic species and 11°00′N; 85°30′–86°00′W; alt. 0–700 m). Although whether any hyperaccumulators of nickel could be sedimentary material covers the northern quarter of the found. In the Caribbean, the rich and largely endemic peninsula, most of the remainder (180 km2) consists of ultramafic flora of Cuba contains the largest number of ultramafic rocks (partially serpentinized peridotites from Ni hyperaccumulators (about 130 species) found any- the Cretaceous) cut by mafic dykes (Fig. 1). The where in the world (Reeves et al., 1996, 1999), mainly Fig. 1. Geology of the Santa Elena Peninsula, Costa Rica (after Tournon, 1994), showing sampling sites: 1. Potrero Grande; 2.Cerro el Inglès; 3. Playa Naranjo. Geological key: 1. Sedimentary cover; 2. Quaternary ignimbrites; 3. Alluvium (a); 4. Dyke swarm, western coast; 5. Ultramafics (partly serpentinized peridotites); 6. Volcanic and sedimentary series, Potrero Grande and southern coast. R.D. Reeves et al. / Journal of Geochemical Exploration 93 (2007) 153–159 155 on soils that have had a very long continuous period of (Near the top of a ridge above the north- exposure and availability for plant colonization, of the western end of Playa Naranjo.) order of 40 million years. The Santa Elena peninsula is Site 3B: Near 10°48.41′N, 85°41.06′W, altitude described as having the oldest continuously exposed 110 m. (Lower on the hillside, near the ultramafic surface in Central America, estimated at edge of the ultramafic soil area.) 85 million years (Janzen, 1986). This raises the possibility of parallels with the Cuban flora in permit- Soil samples and herbarium and analytical plant ting the evolution of significant numbers of endemics specimens were collected from the serpentine sites and Ni hyperaccumulators. Up to the present time, within the areas noted above. The collection consisted of outside Cuba, only isolated additional cases of Ni 73 plant specimens (56 different species, 51 of which hyperaccumulation have been found in the Caribbean have been identified at species level) and 6 soils. The region: Phyllanthus nummularioides in the Dominican main purpose of the soil collection was to confirm by Republic (Reeves et al., 1996) and Psychotria grandis chemical analysis their ultramafic nature, and the degree in Puerto Rico (Reeves, 2003). to which they correspond in composition to other serpentine soils analyzed from many other parts of the 2. Materials and methods world. Soil samples were in all cases taken from the rooting zone of the herbs and small shrubs, i.e. at 0– 2.1. Sample sites and collection 10 cm depth. Herbarium-quality voucher specimens of all plant In this study, sites chosen for investigation were species in this study were collected in duplicate, with the confined to the eastern part of the peninsula, at altitudes number sequence R.D. Reeves 2547–2618 and 2550A. ranging from 100 to 500 m. The serpentine soils here One set has been deposited at the Herbarium of the are brownish–red soils on hilly relief, mostly moder- Instituto Nacional de Biodiversidad, Santo Domingo de ately deep, much eroded, medium- to heavy-textured, Heredia, Costa Rica, and the other set formed part of the with low fertility. Much of the area supports a savanna Massey University Metallophyte Herbarium, Palmer- or cerrado-type vegetation with grasses dominant; there ston North, New Zealand, which has now been placed in are a few trees, scattered shrubs of 1–3 m and a the care of the Herbarium of the Royal Botanic Garden, moderately rich flora of herbs and low (0.5–1 m) shrubs Edinburgh, Scotland. among the grasses. Rainfall is typically about 1500 mm/yr, and strongly seasonal: there is normally 2.2. Analytical procedures less than 100 mm during the 5- to 6-month dry season between November and May. Sampling was carried out The b2 mm fraction of each soil sample was dried for in January 2003. 2 days at 40 °C, ground and sieved through a 70-mesh Collection sites are summarized by the following sieve to give particles b200 μm in diameter. Accurately data, and are indicated in Fig. 1, which shows the extent weighed subsamples of about 0.15–0.20 g were taken of the ultramafics on the peninsula. for analysis. Total element concentrations were deter- mined by digestion with 12 mL of a 1:1 mixture of Site 1: Potrero Grande.